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Vombatidae, Marsupialia) from Miocene Deposits in the Riversleigh World Heritage Area (Queensland, Australia)

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Vombatidae, Marsupialia) from Miocene Deposits in the Riversleigh World Heritage Area (Queensland, Australia) Palaeontologia Electronica palaeo-electronica.org New genus of primitive wombat (Vombatidae, Marsupialia) from Miocene deposits in the Riversleigh World Heritage Area (Queensland, Australia) Philippa Brewer, Michael Archer, Suzanne J. Hand, and Richard Abel ABSTRACT A new genus of primitive wombat, described here, is the most complete early wombat found. Two partial maxillae, as well as isolated teeth, were found within the Riversleigh World Heritage Area in northwestern Queensland, Australia. Previous to this, only a single species of wombat (Rhizophascolonus crowcrofti) was known from deposits older than 10 Ma, despite an estimated divergence date of 40 Ma between wombats and their closest living relative, the koala. We performed a phylogenetic anal- ysis incorporating the new taxon and using dental, cranial and postcranial characters as well as multiple outgroups in order to control for the diverse specialisations of closely related families and the continuing problem of recognising homologies within these groups. The results indicate a sister relationship between the new taxon and R. crowcrofti relative to all other wombats. Based on detailed comparisons between the cheek teeth of wombats, we propose a model which explains the observed differences in cheek tooth form and function in this group. If this model is correct, it could be used to map morphology against diet, and potentially climate. Brewer, Philippa. Department of Earth Sciences, Natural History Museum, London SW7 5BD. [email protected] Archer, Michael. School of Biological, Earth & Environmental Sciences, University of New South Wales, Sydney, 2052, New South Wales, Australia. [email protected] Hand, Suzanne. School of Biological, Earth & Environmental Sciences, University of New South Wales, Sydney 2052, New South Wales, Australia. [email protected] Abel, Richard. Faculty of Medicine, Department of Surgery & Cancer, Imperial College London, London SW7 2AZ, England. [email protected] Keywords: new species; new genus; phylogeny; Marsupialia; Vombatidae; Australia http://zoobank.org/65F19586-5DC7-4DC1-8599-C16743FF024D PE Article Number: 18.1.9A Copyright: Palaeontological Association March 2015 Submission: 28 March 2014. Acceptance: 28 January 2015 Brewer, Philippa, Archer, Michael, Hand, Suzanne J., and Abel, Richard. 2015. New genus of primitive wombat (Vombatidae, Marsupialia) from Miocene deposits in the Riversleigh World Heritage Area (Queensland, Australia). Palaeontologia Electronica 18.1.9A: 1-40. palaeo-electronica.org/content/2015/1060-new-wombat-from-riversleigh BREWER ET AL.: NEW WOMBAT FROM RIVERSLEIGH INTRODUCTION Extant wombats are medium- to large-sized (20-200 kg; size categories 2-3 of Andrews et al., 1979, categories D-G of Hernández Fernández et al., 2006) marsupial herbivores endemic to Austra- lia. There are seven recognised genera, of which two are extant; the Common Wombat (Vombatus ursinus) and the Hairy-nosed Wombats (repre- sented by two species, Lasiorhinus latifrons and Lasiorhinus kreftii). All three of the extant species are burrowing grazers. All wombats are classified in the family Vombatidae and together with six extinct families make up the Infraorder Vombato- morphia (Aplin and Archer, 1987). The closest liv- ing relative of the wombat is the Koala (represented by a single extant species, Phascol- arctos cinereus). Koalas are classified in their own FIGURE 1. Map of Australia showing the location of family, Phascolarctidae, and together with the Miocene wombat fossil sites. Vombatomorphia comprise the Suborder Vombati- formes (Aplin and Archer, 1987). cene deposits, with the greatest diversity of wom- Despite an estimated divergence date bats being found in the Pliocene-Pleistocene. between the Phascolarctidae and the Vombatidae Because extant wombats are specialist grazers, it of 40 Ma (Beck, 2008), only a single monotypic can be hypothesised that their post-Miocene genus of wombat has been described from depos- increase in diversity is related to the spread of its older than 10 Ma. This taxon, Rhizophascolonus more open habitats and greater availability of crowcrofti, was described on the basis of a single graze. However, each of these taxa almost cer- isolated tooth from Lake Ngapakaldi, central Aus- tainly occupied slightly different niches, and the tralia (Figure 1; Stirton et al., 1967), although addi- picture is likely to be more complex. tional specimens have since been described This paper describes specimens of a new (Brewer et al., 2008). genus and species of primitive wombat from early Wombats are significant in being the only Aus- Miocene deposits within the Riversleigh World Her- tralian marsupial group to possess hypselodont itage Area in northwest Queensland (Figure 1). cheek teeth. Hypselodonty is the condition where The specimens comprise two partial maxillae and the teeth grow continuously throughout life. Hyp- isolated teeth, possibly representing as few as two sodonty (high-crowned molars) and hypselodonty individuals. Despite the small sample size, this new (also known as euhypsodonty; Mones, 1982) are taxon represents the most complete early wombat adaptations to high rates of tooth wear and have known. independently evolved in numerous groups. Exam- ples include the cheek teeth of bovids and lago- MATERIALS AND METHODS morphs, respectively. The evolution of hypselodonty in the Vombatidae is poorly docu- Phylogenetic Analysis mented owing to the paucity of fossil material. Of the eight vombatiform families, only vom- Despite this, the Vombatidae are potentially valu- batids and phascolarctids have extant representa- able indicators of changing environmental condi- tives. Interfamilial relationships within the tions in Australia during the Neogene (e.g., Myers Vombatiformes are contentious (Archer, 1984; et al., 2001). Marshall et al., 1990; Munson, 1992; Murray, 1998; The oldest known hypselodont wombat is Black et al., 2012). Accordingly, a phylogenetic Warendja encorensis from the late Miocene of Riv- analysis was performed in which a range of out- ersleigh (Figure 1; Brewer et al., 2007). This genus group taxa were selected (Peramelemorphia and is also found in Pleistocene deposits in South Aus- Didelphimorphia). The choice of outgroup taxa tralia and New South Wales (Flannery and Pledge, concurs with the interordinal relationships deter- 1987; Brewer, 2007), forming a ghost lineage mined by Beck (2008), constrained in some cases throughout the Pliocene. In addition to Warendja, by the availability of postcranial material for some six additional wombat genera are first found in Plio- 2 PALAEO-ELECTRONICA.ORG taxa. A relatively unspecialised representative of at the Museum Victoria; NMV P, palaeontology col- each of the eight vombatiform families was lection at the Museum Victoria, Melbourne; QM F, included in the analysis. A total of 16 taxa were palaeontology collection at the Queensland coded in the analysis. Museum, Brisbane; SAM M, zoology collection at The data set consisted of dental, cranial and the South Australian Museum, Adelaide; SAM P, postcranial characters taken from the literature and palaeontology collection at the South Australian personal observations. Where characters were Museum, Adelaide; AR, temporary number for taken from the literature, they were re-scored, palaeontology collection at the University of New where possible, for each of the taxa in the analysis South Wales, Sydney; UNSW Z, and UNSW J, in order to verify the results of others and to ensure zoology collection at the University of New South consistency. Some characters used by other Wales, Sydney. authors in their phylogenetic analyses were rejected from this analysis because they were SYSTEMATIC PALAEONTOLOGY found to be highly intraspecifically variable (e.g., Order DIPROTODONTIA Owen, 1866 presence of frontal-squamosal contact: Springer et Suborder VOMBATIFORMES Woodburne, 1984 al., 1997; Wroe et al., 1998; Horovitz and Sánchez- Family VOMBATIDAE Burnett, 1830 Villagra, 2003; Black et al., 2012), or they were not Genus NIMBAVOMBATUS gen. nov. parsimony-informative for any of the taxa exam- ined. zoobank.org/C832B075-FB33-4C41-B5DF-EA48888AA8D3 A total of 115 characters were used in the Etymology. ‘Nimba’ is the Waanyi (the local analysis of which 28 are of the type ordered (Wag- Aboriginal language of the Boodjamulla region) ner) and 87 are of the type unordered. The data word for ‘small’. ‘Vombatus’ means ‘wombat-like’. matrix was analysed using Paup 4.0b10 (Swofford, Diagnosis. As for the species diagnosis until addi- 2002) using PaupUp graphical interface (Calendini tional species are described. and Martin, 2005). The branch and bound search Nimbavombatus boodjamullensis sp. nov. option was utilised, using a mixture of ordered and Figures 2-5 unordered characters (Appendices 1, 2). Multistate characters were treated as polymorphisms. All zoobank.org/1D7CD6A8-DB90-4D89-9B2F-7306C2DFDAA5 characters were presumed to have equal weight. Etymology of Specific Name. Refers to the Accelerated transformation (ACCTRAN) and Waanyi (the local Aboriginal language of River- delayed transformation (DELTRAN) character- sleigh) word for the Riversleigh area (Boodjamu- state optimisation methods were employed. Boot- lla). strap support (50%) for clades was determined Holotype. QM F23774 (left maxilla with C1, P3 and using 10,000 replicates. M1-4) Paratypes. QM F23725 (left maxilla fragment with Terminology P3, M1-3 and partial alveolus of M4); QM F30339 Incisor, premolar and molar homology follows 1 (right
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